Disequilibrium in two-locus mutation-selection balance models.

Equilibrium behavior of two-locus mutation-selection balance models is analyzed using perturbation techniques. The classical result of Haldane for one locus is shown to carry over to two loci, if fitnesses are replaced by marginal fitnesses. If the fitness of the double heterozygote is smaller than would be produced by a multiplicative model, as in additive or quantitative fitness models, the disequilibrium is negative--an excess of gametes with one rare allele. In this case the disequilibrium can be as large as one-half its maximum value possible, if the recombination rate is small, not greater than the strength of selection. If the fitness of the double heterozygote is larger than would be produced by a multiplicative model, the disequilibrium is positive, and is very small relative to its maximum value possible, even if the recombination rate is zero.